Vocal variability of chaffinch song +

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Vocal variability of chaffinch song
(
Fringilla coelebs L.)
Astakhov
a
O.A. (dissertation) 2006.
A general characteristic
•
In
early
works
geographical
variability
of
song
at
many
species
of
birds
has
been
marked
as
a
phenomenon
of
differentiation
of
song
structures
or
song
patterns
(as
the
image
on
sonograms
)
in
different
local
populations
(Marler,
1952
;
Marler
and
Tamura,
1962
;
Sick,
1939
;
Poulsen,
1951
)
.
•
Local
variants
of
vocalizations
of
birds
can
be
considered
as
analogy
of
human
speech
and
to
name
as
dialects
.
The
word
"dialect"
has
taken
place
from
linguists
and
means
a
local
speech
variant,
characteristic
for
region
.
Borders
of
a
dialect
define
lexical
(
word
structure),
morphological
(an
accent,
structure
)
and
phonetic
(pronunciation)
characteristic
s
of
the
dictionary
(Kurath,
1972
)
.
•
Song
dialect
of
birds
is
a
variant
of
the
traditional
form
of
song,
shared
by
members
of
a
local
population
of
birds
and
forming
borders
of
a
dialect
(separating
from
other
variant
of
song
pattern),
within
the
limits
of
which
there
is
a
traditional
training
(song
learning)
of
characteristic
song
components
of
the
given
population
.
But
thus
big
song
repertoir
e
s
of
some
species
of
sparrow
birds
interfere
(have
problem)
with
objective
definition
of
dialect
borders
(Thielcke,
1969
;
Kreutzer,
1974
;
Kroodsma,
1974
;
Baker,
1975
;
Baptista,
1975
;
Lemon,
1975
;
Payne,
1981
;
Mundinger,
1980
,
1982
)
.
•
Chaffinch
(
Fringilla
coelebs
L
.
)
is
classical
object
of
study
of
becoming
of
vocal
repertoir
e
(Thorpe,
1958
;
Marler,
1956
;
Nottebohm,
1967
)
and
of
geographical
variability
of
species
specific
song
in
a
population
(
Promptov
,
1930
;
Sick,
1939
;
Thielcke,
1961
;
Simkin,
1983
;
Slater
e
t
al
.
,
1984
)
.
One
of
problems
of
our
researches
has
consisted
in
revealing
macrogeographical
distinctions
of
chaffinch
song
(
Fringilla
coelebs
L
.
)
in
different
local
populations
,
which
are
removed
approximately
on
1000
km
from
each
other
.
In
northwest
(
Curonian
spit,
the
Kaliningrad
region
)
and
in
central
parts
(
Zvenigorod
,
Moscow,
Michurinsk)
of
the
European
Russia
(fig
.
1
)
have
been
made
tape
records
of
singing
males
(N=
218
)during
the
spring
-
and
-
summer
period
of
2005
-
2006
.
Fig. 1
.
Map of Russia (points mark places of sampling of songs)
Methods of research
•
Regional variability
of song can be determined in qualitative aspect (the form of a syllable, syntax) and quantitative (time
-
and
-
frequency) parameters. The concept of a dialect should be closely (cautiously) and is carefully argued in an establishment of regional variability call and song
patterns.
•
At the analysis
of
song sonogramms
basically applied two qualitative methods
(Mundinger, 1982):
1. Revealing of phonetic distinctions
(frequency of a sound, its form on sonogram
) or of ways of a pronunciation of syllables of the phrases making songs;
2. Revealing of lexical distinctions
(changes of phrases of songs as a whole). •
A
lso have been spent the quantitative analysis
of dialect forms of songs of one type and comparison of their basic time
-
and
-
frequency parameters
(duration of songs
-
sec
, number of elements in type of song, duration of syllables in a trill, the maximal, minimal and average frequency of song
-
KHz
, intervals between songs
-
sec
).
•
Sonograms of
types of songs have been analyzed with the help of computer program Avisoft SASLab Light
. In total have been analysed about five thousand songs. Types of songs have been
marke
d by L
atin letters
. At record, songs of one type have been met in different points of territory
(considered that belong to repertoires of different males), therefore alongside with the letter have been designated by numbers in ascending order (for example, А
1, А
2, А
3, etc.).
Results and discussion
•
In populations of the central part
of the European Russia (N=65
of males
) we allocate (
distinguish)
15 types of song
of chaffinch
(
Fringilla coelebs L.)
,
which have been completely similar in structure or have been in part modified in the syllabic form (on sonograms
) in comparison with songs of same
types
in samples of a northwest part
of the European Russia (N=153
of males
). •
For seven types of songs
from twenty two (tests on Curonian spit
) analogies (similarities) ha
ve
not come to light –
probably, owing to their rarity
. Many song
types in the central part of the European Russia have been considered
as combined phrases
(
parts) of syllabic patterns known to us, but frequently with the changed figure
(forms)
on sonogram
.
•
In result
,
have been found 12 dialect forms of songs of
one type
,
which appeared similar in base structure
of elements, of phrases, but in different regions of Russia frequently had
distinguished manners, ways of their performance at singing
(phonetic aspect).
•
The average size
of chaffinch repertoir
e
(
Fringilla coelebs L
.
) in populations of the central part
of the European Russia from statistical calculations has been submitted 1,93 ±
0,22
type of songs (max –
4 types of song, min –
1 song type) at a capture (volume) on the average 24,7 ±
11,3 songs from one of male
.
•
The basic time
-
and
-
frequency parameters
of local variants of types of songs
of chaffinch
(Fringilla coelebs L.)
at the quantitative analysis had
insignificant macrogeographical variability
that can speak about their importance in transfer
and maintenance species specific song
traditions
. The most stable parameters of local variants of songs of one type
appeared minimal and average
(median) frequency
, values of length
and intervals of elements of a trill
.
The table.
Character of vocal variability of chaffinch songs (Fringilla coelebs L
.)
in populations of northwest and the center of the European part of Russia (N = 218) *
Vocal variability
Song type (
token
) A
B
C
D
E
F
G
I H
J
K
L
M
N
O
P
Q
R
S
T
V
W
U
Total
Dialects
1
+
+
+
+
+
5
Subdialects
2
+
+
+
+
+
+
+
7
European part of Russia
Sample size of song type (n) in population
Northwest
18
5
38
11
7
16
10
22
6
13
3
1
20
8
18
1
1
24
17
14
2
8
183
Center
8
8
24
3
5
1
11
12
2
5
2
2
3
3
93
Note:
1
dialects –
different phonetic norms of song types; 2
subdialects –
small phonetic distinctions of song types; * -
sample size of chaffinch males (Fringilla coelebs L
.)
at record; the most widespread (frequently met) in populations types of songs (with sample n> 15) are allocated (distinguish) by a font.
Probably, at song types are found
the dialects (vocal variability), because of their high frequency of occurrence in populations at record. Perhaps, other types of song also have dialects in this territory, but have not come in our sample.
Thus, on twelve types of chaffinch songs
(Fringilla coelebs L
.)
the dialect forms
(B C D F G I J M N V S U) have been found. It is interesting to note, that dialect forms of songs or separate phrases (in the "combined" songs) of one type can meet within the limits of one local population. Proceeding from samples, made by us in different regions
of the European Russia (
northwest
and the central part
), it is possible to draw a conclusion, that on some song types of chaffinch (Fringilla coelebs L
.)
exist subdialects
(insignificant phonetic distinctions of syllables) -
for example, on song types B, M, D, G, V, S, U, and dialects
(different phonetic norms of song types) -
C, C * and C
#
, F, I, J, N, and are also stable song types (in samples of different regions unchanged) –
song type A.
As
a
bright
example
of
vocal
variability
of
chaffinch
(Fringilla
coelebs
L
.
)
it
is
possible
to
result
(to
show)
some
local
variants
of
songs
,
which
we
have
attributed
to
one
type
(fig
.
2
,
3
,
4
)
.
Samples
of
types
of
song
,
which
will
be
resulted
(be
shown)
,
were
not
single
in
a
population
and
have
been
recorded
in
repertoir
e
s
more,
than
1
-
2
individuals
.
The song
t
ype V
1
2 Fig. 2
.
Dialect forms
of
song type V: 1 –
the song
type V 1 (record on Curonian spit
, the Kaliningrad region); 2 –
the song type V (record in Zvenigorod
, the Moscow region) Fig
. 3
.
Dialect forms of song type
S: 1 –
the song
type S5 (record on Curonian spit
, the Kaliningrad region); 2 –
the song
type S (record in Zvenigorod
, the Moscow region).
The song t
ype
S
The song
t
ype
I
Fig. 4
.
Dialect forms of song type
I: 1 -
the
song
type I 6 (record on Curonian spit
, the Kaliningrad region); 2 -
the song
type
I 1
2
(record in Moscow); 3 -
the song
type
I 6 (record in Zvenigorod
, the Moscow region).
1
2
1
2
3
Despite of stability in a manner of performance of song type B, in different local
populations
, on sonogramms
it is possible to note some variability of the
final stroke
(in the latter case (fig. 5.4) the number of the first elements of a stroke is increased, and the phrase is heard as a trill). The
s
ong
t
ype
B
Fig. 5
.
Local variants of song type B (from different areas of Russia): 1 –
the song
type В5 (record on Curonian spit
, the Kaliningrad region); 2 –
the song type B
(record in Zvenigorod
, the Moscow region); 3 –
the song
type В8 and 4 –
the song
type В7 (record in Michurinsk, the Tambov region).
1
2
3
4
At the quantitative analysis of local variants
of song type B (with the help of computer program Avisoft
-
SASLab Light
) come to light some distinctions
(differences)
in values of the basic time
-
and
-
frequency parameters, but in the majority they are similar (tab. 1).
Table 1.
The basic time
-
and
-
frequency parameters of local variants of song type B
The note:
average value and a standard deviation of parameters of song types from statistical calculations for all songs of one type which were reproduced chaffinch males in the given points of record are specified; the strongest differences counted a difference of parameters
>0,5
KHz
in frequency
and >0,02 sec
in length (are allocated (
distinguished)
by a font); * -
the trill of
song type
B
will consist of two phrases (fig. 5).
The length of songs, number of their syllables
(elements)
, the maximal frequency, intervals between songs and length of elements of the second phrase of a trill have the values varying in wide enough range. The minimal frequency
and length of syllables of the first phrase
of a trill
of song
type B are appeared completely similar
as in northwest (
Curonian spit
), and in the center of the European Russia (Zvenigorod
, Michurinsk). Type of song (token
)
The name of record
place
The length of song
, sec
Min frequency
, К
Hz
Max frequency
, К
Hz
Median
(average) frequency, К
Hz
Number of syllables in song
type
Length of syllables (trill)
, sec*
Intervals between songs
,
sec
1
phrase
2
phrase
В5
Curonian spit
2,47
±
0,104
1,57
±
0,14
7,19
±
0,32
3,77
±
0,14
24,5
±
0,93
0,043
±
0,003
0,14
±
0,01
5,32
±
1,9
В
Zvenigorod
(
Moscow
region
) 2,48
±
0,36
1,68
±
0,16
7,92
±
0,31
3,83
±
0,36
22,25
±
2,99
0,04
±
0,003
0,17
±
0,008
В8
Michurinsk (
Tambov regio
n)
2,34
±
0,15
1,62
±
0,09
8,044
±
0,53
4,12
±
0,29
22,3
±
2,6
0,034
±
0,006
0,127
±
0,013
4,3
±
0,59
В7
Michurinsk (
Tambov regio
n)
2,99
±
0,11
1,67
±
0,08
7,63
±
0,08
3,79
±
0,18
27,4
±
1,17
0,033
±
0,003
0,15
±
0,007
7,06
±
0,19
In samples types of songs have been found
, which are difficult for differentiating among themselves because of their similarity (fig. 6).
Fig. 6
. D
ifficultly differing chaffinch song types (record on Curonian spit
, the Kaliningrad region): 1 –
the
song type D3; 2 –
the song type F4; 3 -
the song type G3. We have attributed the given samples of songs to different song types (D, F, G), but nevertheless they are similar
among themselves in base structure of elements
(especially in the started singing
and trill)
,
which nevertheless are performance
at singing by different manners, ways
that makes their distinct
or different
. Probably, it –
the initial forms of the further modification
,
being in themselves insignificantly transformed forms for precise revealing of type to which are attributed.
1
2
3
Many types of chaffinch songs (Fringilla coelebs L
.)
in samples of the central part of the European Russia were considered by us as combined phrases
(parts)
of already known types of the songs which have been recoded on Curonian spit (
the Kaliningad
region), but their elements have been modified
on sonograms
. It is possible to result some examples.
Fig. 7
.
Dialect forms of phrases from song types B and F: 1 –
the song type B (record on Curonian spit
, the Kaliningrad region); 2 –
the combined song type ВF 5 (record in Michurinsk, the Tambov region); 3 –
the song type F (record on Curonian spit
, the Kaliningrad region)
The song type ВF 5 (fig. 7.2) is recoded in the central part of the European Russia (Michurinsk) and will consist of the modified combined phrases: the trill is similar with elements of started singing
of song
type F (fig. 7.3); the final stroke also is similar to song type F which has been recoded on
Curonian spit
; and first two phrases of song
type ВF5
-
from
song
type B
(fig. 7.1). Modification of forms of elements of one type can speak about their dialects (
special to a manner of a pronunciation) in a local population
. 1
2
3
Table 2.
The quantitative analysis of dialect phrases of song types B and F
Type of song
(
place of record
)
Parameters of
phrases
В
(
Curonian spit
)
В
F
5
(
Michurinsk)
F
(
Curonian spit
)
1
phrase
(
started singing
)
2
phrase
(
trill
)
В
-
similar
elements (
started singing
)
F
-
similar
elements
(
trill
)
1 phrase
(
started singing
)
1
phrase
2
phrase
The length of phrase
, sec
1,13
±
0,09
0,32
±
0,05
0,62
±
0,13
0,24
±
0,02
1,17
±
0,25
1,27
±
0,11
Max frequency
, К
Hz
6,57
±
0,46
4,626
±
0,32
7,3
±
0,26
5,9
±
0,52
7,44
±
0,22
6,25
±
0,2
Min frequency
, К
Hz
2,017
±
0,13
2,067
±
0,1
3,1
±
0,3
2,4
±
0,12
1,9
±
0,17
1,722
±
0,14
Median (average) frequency
, К
Hz
4,306
±
0,24
3,17
±
0,09
5,44
±
0,2
3,8
±
0,17
4,31
±
0,172
4,2
±
0,09
Number of syllables in phrase type
11,43
±
0,79
5,29
±
0,76
5,2
±
0,8
5
±
0
6,4
±
1,34
7,6
±
0,7
The length of syllables
, sec
0,058
±
0,01
0,037
±
0,005
0,08
±
0,014
0,04
±
0,008
0,15
±
0,01
0,12
±
0,028
Intervals between syllables
,
sec
0,048
±
0,01
0,037
±
0,004
0,05
±
0,01
0,02
±
0,005
0,05
±
0,008
0,06
±
0,012
The note: average value and a standard deviation of parameters of dialect phrases from statistical calculations for all songs of one type which were reproduced chaffinch males in the given points of record are specified; the strongest differences of phrases of one type counted a difference of parameters
>0,5 KHz
in frequency and >0,02 sec
in length (are allocated
(
distinguished)
by a font).
Proceeding from the analysis (tab. 2) carried out by us, it is possible to draw a conclusion, that the length of dialect phrases
and their maximal frequency
basically are various
, and the minimal frequency
and intervals between syllables
appeared are similar
in the majority. Values of others frequency
-
time parameters can to vary widely
, but within the limits of the certain range.
It is interesting to note, that at the greater distinction of the form of syllables
(elements)
of dialect phrases on sonograms
(
up to occurrence of new syllabic patterns)
are observed
the big distinctions
of their basic frequency
-
time parameters
(for example, the first phrases of the started singing of song types B
and BF). Fig. 8.
Dialect forms of phrases of song types Т and S: 1 –
the song type Т (record on Curonian spit
, the Kaliningrad region); 2 –
the combined song type ТS (record in Michurinsk, the Tambov region);
3 -
the song type S (record on Curonian spit
, the Kaliningrad region) +
Thus, in samples of different areas
of Russia there can be the song types combined from phrases
of
song patterns of
other populations
, but with the modified elements
(syllables) –
it is possible to tell, their
dialect forms.
1
2
3
Conclusions
•
Many characteristics of singing of birds are precisely mentioned by social training traditions
, receiving their vocal patterns by species specific imitation
. Therefore the tradition, custom should be taken into account at the analysis of geographical variability of vocal behaviour
. •
In local populations
are formed certain song cultures
which are capable to
change
du
ring time
and can to make dialect forms
on all area of distribution of a species
. Steady song dialects during time –
the phenomenon of conservatism
of the vocal traditions, transmitted to the
subsequent generations
by means of vocal imitation. •
For many species
the
patterns of variability
as purchases
(changes)
of vocal traditions are a product
(result) of cultural evolution (Mundinger, 1980). Cultural evolution
is a parameter for species
with a historical variety of patterns of microgeographical variability, and also for species
with differentiated
syllables
. But it is not a parameter in morphological variability (Lemon, 1975; Slater and Ince, 1979).
•
Vocal norms
(installation
s
), likely, can be judged by quantity of individuals in a population, which adher
e
to those or others song
cultures
(ways of singing)
of different
song types. If the majority of
males
sing any song
type such certain style
(
special manner of performance), then it
is
will be vocal norm
(installation) i
n the given song
type (or in
type of a phrase) in the
given population
. But it is difficult to define vocal norms of a population at repertoir
e
s of great volume
(when
song types
much) (Kroodsma, 1974). •
For
chaffinch
(Fringilla coelebs L
.)
typically selective training (song learning) of species specific song
in sensitive
period (Marler, 1956; Thorpe, 1958; Nottebohm, 19
6
7). Therefore young birds do not learn songs of other species
, and are guided by a time
-
and
-
frequency range of songs of their species
. Probably, as a result of such rigid genetic determination
of species species
song (Simkin, 1983) is observed small variability
of the basic time
-
and
-
frequency
parameters of its different dialect forms
(even at the account of a wide variation of values).
•
Proceeding from samples
, made by us in different regions of the European Russia
(northwest and the central part), it is possible to draw a conclusion, that i
n some
song types of
chaffinch
(Fringilla coelebs L
.)
exist subdialects
(insignificant phonetic distinctions of syllables) –
for example, i
n song type
s
B
, M, D, G, V, S, U, and dialects
(different phonetic norms of types of song) –
C
, C
#
and C
*, F, I,
J, N, and there are also stable types of song
(in samples of different regions are invariable
) –
song type A.
•
By f
ull dialect
of chaffinch song (Fringilla coelebs L.)
in the certain territory it would be more correct to ac
count the presence of dialect forms on all
song
types
known to us in a population (originally
22
types
have been found), that, likely, it is almost impossible
because of limitation of a sample
and constant mixing of vocal traditions
of birds in
different populations as a result of
migrations
(Slater and Ince, 1979, 1980; Espmark et al
.
, 1989).
The literature
•
Promptov
A
.
N
.
1930
.
Geographical
variability
of
chaffinch
song
in
connection
with
the
common
questions
of
seasonal
flights
of
birds
//
Zool
.
journ
.
№
10
(
3
)
:
17
-
40
.
•
Simkin
G
.
N
.
1972
.
About
biological
value
of
bird
singing
//
The
Bulletin
of
the
Moscow
university
ю
№
1
.
Moscow
:
34
-
43
.
•
Simkin
G
.
N
.
1983
.
The
typological
organization
and
population
phylogeny
of
bird’s
songs
//
Bulletin
of
Moscow
community
investigators
of
nature
.
Section
of
biology
.
V
.
88
.
№
1
.
Moscow
:
15
-
27
.
•
Baker
M
.
C
.
1975
.
Song
dialects
and
genetic
differences
in
White
-
crowned
Sparrows
(
Zonotrichia
leucophrys
)
//
Evolution,
29
:
226
-
241
.
•
Baptista
L
.
F
.
1975
.
Song
dialects
and
demes
in
sedentary
populations
of
the
White
-
crowned
Sparrow
(
Zonotrihcia
leucophrys
nuttalli
)
.
-
Univ
.
Calif
.
,
Berkeley,
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